title population genetical study of natural hybridization ... · title population genetical study...
TRANSCRIPT
Title Population genetical Study of natural hybridization betweenPapio anubis and P. hamadryas( Dissertation_全文 )
Author(s) Shotake, Takayoshi
Citation Kyoto University (京都大学)
Issue Date 1981-03-23
URL http://dx.doi.org/10.14989/doctor.r4378
Right
Type Thesis or Dissertation
Textversion author
Kyoto University
影} 聯馨
㌔,∴ 、べ ぞ罐_蘇 ・
牟擁 灘
」
Pρpulati。ngene七icals七udy・fnaturalhybridi・ati。nbetween
迦anubi。andR.h。m。d。ya。')
・虹
凸
京大 附図
TAKAYOSH工SHOTAKE
KyOtOUniVerSity
ABSTRACT.Thena『 ヒuraユhybridizationbetween .堕anubisand
王1.,hamadryasincentralEthiopiawasstudiedfromapopulation
geneticaユperspective・Studiesweremadeusingelectrophoreticaユ
bloodproteinvariationsasmarkersinordertoclarifythe
geneticrelationshipbetweenthem.Atotalof563samplesfrom '10popula'ヒionswhichwerecollecrヒedinthefieldstudieswith
asocioecologistinl976andl979wereexaminedfor34blood
proteinloci.Tlenofthe3410cishowedpolymorphism.TheTf,
PA-2andEswerefoundtobeeffectivefordiscriminatingbetween
theanubisandthehamadryas.Geneticvariability,hybridiza七ion
rate,qeneticdistance,migrarヒionratesandcorrelationsbetween
geneticalandmorphologicalandbetweengeneticalandbehavioral
indiceswerecomputedandanalyzed.Theresultsofthepresent
geneticsurveyrevealedthatmostofthepopulationsfromwhich
theauthorcollectedbloodsamplesweremoreorlesshybridized・
TheNeils(1975)qeneticdis七ancebetweenthetwospecieswas
estimatedtobeO.0679atmos'ヒ.Asthisvaluesistoos皿allto
considerthesespeciesasrealbioloqicalspecies,itissupposed
thatthenaturalhybridzoneisfairlywideandstillexapnding
nOW.
1) Thisworkwassuppor七edinpartbyGrantsinaidforscientific
research(Overseasscientificresearch,1975)andofthe
O▽erseasSpecialResearchProqra㎜eofthePrimateResearch
工nstitute,Kyo七 〇Universityinl978byMinistryofEducation,
ロSc■enceandCulture.
一1一
工NTRODUCT工ON コTaxonomyofthegenusPapiohasbeenユnacontroversyover
thepast20years(Tappenl9603De▽oreandWashburnl9633
Buettner-Januschl9631Jollyl9663Rothl9653Grovesl9723
Maplesl9723JollyandBrettl9733SzaユayandDelsonユ9793
工wamo・ ヒoユ980).Thenaturalhybridizationbetweenthehamadryas
baboon,旦 些 ≧amadryasξLndtheanubisbaboon'王 盗 ≧主≦≧anubis'
cohsideredtobeclearlydifferen「 ヒspeciestaxonomically,wasdis-
coveredbyKummeretal。(1970)andNagel(1973).工nl975andl978,
Dr-MasaoKawaisentscientificexpeditionstoEt二hiopiatostudy
thisnaturalhybridizationphenomenonsocioecologically,morpho-
loqicaユlyandgene・ ヒically.Theauthortookpartintheseexpedi-
tionstoanalyzethepresentgeneticstatusofthehybrids.He
madefieldsur▽eysincentralEthiopiainthevicinityofthe
AwashNationalParkwi七hDr.K.Sugawara,socioecologist,under
thesupervisionofDr.Kawaitogeヒgeneticdataoveraperiod
oflOmonths.Proteinelectrophoresisisthemostefficient
techniqueforquantifyinggeneticdifferencesbetweenclosely
relatedspecies(Selanderl976)becauseof七hese工ectiveneutrali-
tyofproteinvariations(Kimural9683Neil974,1975).The
authorhadpreviouslystudiedthepopulationgeneticsofJapanese
macaque,1唖acaca£usga』 皇,andhadundertakenasystematicstudyof
thegenusMacacausingbloodproteinvariatiorlsasageneticmarker
(Shotakel974;Shotakel9793ShotakeandNozawal9743Shotake,
OhkuraandNozawal9753Nozawa,ShotakeandOhkuraユ975,Nozawaet
al.1975,1977,inpress).Hefurtherintendedtostudythe
populationgeneticsand七axonomyofthebaboonsbyusingthesame
method(Sho七ake,NozawaandTanabeユ977,Shotakel980).
一2一
Sofar,someqeneticvaria・ ヒionsofbloodproteininthe
qenusPapiohavebeenrepor'ヒed(Buettner-Januschl9633Barnicot,
EuehnsandDancel9653Barnicot,JollyandEadel9673Ki'ヒchin,
BarnicotandJollyl9673McDermid,VosandDowningl973301ivier,
Bue七tner-JanuschandBuettner-Januschl9743CrawfordandO「Rourke
l9783Lucottel979).However,mostofthesestudiesdidnot
attempt'ヒomakequan・ ヒitativecomparisonbetweehpopulationsor
species,whereasthetaxonomic3tatusofthespeciesin七hegenus
ロ ロ ロコPapユowas■naconrヒroversy・Theauthorpublishedaprel■mユnary
paperdealinqwiththebloodpro七einpolymorphismsin堕
anubis,」 ≧些hamadryasand'theirhybrid,andwithestaimtesof
thea皿ountofgeneexchangeandthegeneticdis『 ヒancebetweenthe
twospeciesfromthedataofthefirstexpeditioninl976
(Sho七ake,NozawaandTanabe,1977).Thepresentpaperisan
extensionwhichpresentsadditionaldataandresultsohヒainedin
thesecondexpedi七ioninl979andafewcorrectionsofpreliminary
repor『 ヒ・
MAT'ER工A丑SANDMETHODS
Thefieldsurveyandbloodsamplingwerecarriedoutin
bo七hl976andl979atthespecies-borderbetweentheanubisand
hamadryasbaboonsintheAwash▽alleyandonthenorthernedge
ofAwashNationalParkwherehamadryasbaboonslivedsouthof
CassemRiver-Nagel(1973),KawaiandSugawara(1976a,b),
Suqawara(1979,inpress)giveade・ ヒailedinterpre'ヒationonthe
hybridzonebetweentheanubisandhamadryasbaboons.InFig.l
andTableltheplaceswherebloodsamplingwascarriedoutare
indica七edbyalphabeticalabbreviation.
A工:1…heanubisbaboonsof工to'ヒ ヱooP.
Theanubisbaboonswhichhadasleepingsiteingallery
一3一
forestlocated5kmabovetheAwashFalls(Fig.1》.Thistroop
consistedof73individualsincluding2hamadryasadul七males
whichhadi㎜igratedintothistroopin1976・However,inl979
thesizeofthistエoophaddecreasedto53individualsincluding
onehalnadryasadultmalewhowasdifferen七from2individuals
observedinl976.Atotaユof27bloodsampleswereobtainedin
l976andl979.
AW=theanubisbaboonsof.We■enchi土 ゴ..troop-swhichlivedinsouth一一、Thirteenbloodsampleswerecollectedfromtheanimals
whichhadbeencapturedbyanimaldealerslivinqinHodjoTown
nearAddisAbabainl976.Theplacewheretheseanimalswere
capturedwaslocatedaboutl5kmsouth-・westGaribaldiPass,
whereDr-Kummer(1968)sawahybrid皿aleandseveralhybrid
femalesduringhisl960survey.
AK:theanubisbaboonsofKenya
TheseconsistedoftheindividualsimportedbytheJapan
塾40nkeyCentrefromKenya20yearsagoandtheirdescendants.
Theauthortook28blQodsamplesfromtheseanimalsbefore
leavinqforl975expeditionandhadstoredtheminadeepfreezer
at-900C.
MG3thehybridbaboonsofGorgetエoQp
Ahybridbaboontroophadasleepinqsiteonacliffa七4
kmbelowtheAwashFalls(Fiq-1).Thistroopwasintensiveユy
studiedbythesociobiologis七 〇four幽parセyandwasfoundto
consistof63individualsinl976(Sugawara,1979).Theauthor
gotl8bloodsamplesandsomemorphologicaldatafromthis
troopin1976・
M1く3thehybridbaboonsofIくerrayutroop
一4一
Atypica工hybridbaboontroophadsomesleepingsitesona
cliffataboutユOkmbelowtheAwashFallsnearKerrayulodge
(Fiq.1).Thistroopwhichwasalsostudiedintensivelybythe
sociobiologistandfoundtoconsistof85individuals(Suqawara,
inpress).Theau七horgot43bloodsamplesandsomemorpholo9-
icaldatafromthistroopinl979.
HBAandHBB=thehamadryasbaboonsofBassakaAandBbands
Thesebandssome七imesjoinedtoqetherandmadeatroQPin
asleepingsiteユoca七edonthenorthernedqeofLakeBassakaat
thefootofMt.Fantale.Atfirst,i七wasthoughtthatthis
troopswasofpurehamadryasbaboons,butafterhabituation,it
wasfoundthatthetroopincludedtwoadultanubismalesand
sevenhybridmales・Thirty-ninebloodsampleswerecollected
fromtheAband(70individuals)andl9fromtheBband(45
individuals)inユ976.
HS3thehamadryasbaboonsofSabureGorqetroop
Thirty-fivebloodsampleswerecollectedfromanimals
wh車chhadbeencapturednearSaburevillageandtransferredto
Metaharabyananimaldealerinl979。
HC3thehamadrasbaboonsofCassemGoretroops
Forty-ninebloodsampleswereobtainedfromtheanimals
whichhadbeencapturedalonqtheCassemRiverandtransferred
toMetaharabyananimaldealerinl979・
HAG:thehamadryasbaboonsofAwashGameReservetroops
Thesesampleswerecollecteda『 ヒthestockfarmofanimal
dealers,whocaughttheanimalsfromseveral七roopsユivinqin
AwashGameReservelocated70㎞nor七heastfromAwashStation.
Thiswasonthenorthernlimitofthehybridzonedescribedby
Nagel(1973)(Fig・1》 ・Bloodsamplingwascarriedoutseveraltimes
一5一
ineachpopulationfoτabou七twomon七hsfromJanuarythrough・
March,1976andforabou七5mon七hsfromFebruarythroughJuly,
1979.Thesampleswerebrought七 〇 七heNAMRU5工nstitu七ein
AddisAbabainl976andtoapriva七e工talianlaboratoryin
AddisAbabainl979.Redcellsandplasmaweresrヒored
separatelyat-60。Cinl976andat-25。Cinl979un七ilthe
author冒sdepaτturetoJapan.
Thirty-fourgeneticlocicontrolling31kirldsofblood
pro七einswereexaminedbystarchandagarqelelec七rophoreses.
TheyarelistedinTable2。Samplingofapro七einlocuswas
random'上}avinqnorelationtowhethertheexistenceofgenetic
variationatthislocuswaslikelyornot.Forsomeproteins
thepresenceofageneticpolymorphismandthemodeof
inheritanceinthegenusPapiohavealreadybeepdetermined
【forTf,seeButtner-Janusch(1963),Barnicot,etal.
(1965)andNcDermid,VosandDowning(1973)3forCA一 工,see
Barnicot'etal・(1965)`andBarnicot,JolユyandEade・
(1967)andOlivier,Bue七tner-JanuschandBue¶ ヒtner-Janusch(1974)3
for6PGDandPi,seeMcDermid,VosandDowninq(1973)].The
sys七emofsymbolsforpolymorphiclociusedinthispaperdiffers
fromthatadoptedbytheaboveauthorsonaccountofdifficulty
ofidentification.AsforthemOdeofinheritanceofother
variableproteins,codominancerelationswereassumedbecause
equivalentelectrophoreticbandswereobservedinpostulated
herヒerozyqotes,A七allgene七icloci,theallelefrequenciesin
individualgroupscouldbecalcula'ヒedbyasi皿plegene-coun七ing
merヒhod.
Thegeneticvariabilitywithinpppulationswasquantified
bymea・u「ingthep「opo「tionofp。1顕o「phicl。ci(Pp。 ・y)(the
cri七erionofpolymorphismwasthefrequencyoftheco㎜ones七
一6一
aユleles<0.99)3theexpec七edproportionofheterozygotesper
individual(H=1一 ζq呈whereqiw・ ・th・frequen・y・fi-th
alleleatalocusandtheaveraqewasoveralllociincludinq
lociwirヒhoutvariation)3andtheeffectivenu㎡berofalleles
pe「1・cu・(程=1/苧qi)・
Fromthegenefrequencydata,thegeneticdistancewas
calculatedineverypairofthepopula七ions.Thegenetic
distancewasob七ainedfromthenormalizedidentityofqenesas
devisedbyNei(1974,1975).Thatis,thedisrヒancebetweenthe
」騨七handk一 七hp。pulati。n'Dゴk・waヤca'cu'atedasDjk='1。9・ エ'
whereエ ー ζqi」qik/(ζqil● ζqi灸)7・ エnth・f… ・ae・w・ ・th・
n。 「malizedidentity・fgene・(qen・ticsimila「ity)'qijandqik
werethefrequenciesof七hei-thallelesatalocusin七hej-th
andk-thpopulations,respectively,andtheaverageswereover
allthegenelociexaminedincludinglociwithoutvariation.
RESULTS
Descriptionofbloodpro七einolorhism
Ofthe34qeneticlociexa皿inedlOloci,thatis,Tf,Pi,
PA-2,PlasmaEs,6PGD,PGM一 工,PGM一 工工,ADA,CA-IandAKshowed
polymorphism.工nthepreviousreport(Shotake,Nozawaand
Tanabe,1977)TBPAlocuswaspolymorphic,butitwasclarified
bythesucceedingexaminarヒionthatthislocuswascompletely
monomorphic・Thislocuswasnotthereforeusedfoτqenetic
analysisinthepresentwork.Theauthorfoundanewallelein
pA-210cusinthepresentstudy.Thepatternsoftheelectro-
phoreticvaria『 ヒionsa∫eshowninFig・2・Table3givesthe
frequenciesofdifferentallelesoccurrinqa・ ヒeachlocusinthe
lOpopulationsofbaboons.Detailedin・ ヒerpre・ ヒationsofthese
variablelociareasfollows.
一7一
Tf(Plasmatransferrin)=
Theauthorfoundeightkindsofqenotypes,thatisDD,DD冒,
D璽D㌧D冒E,DE,EE,DICandECcontrolledbyfourallelesatthis
locusamongthehamadryasbaboons,theanubisbaboonsandtheir
hybrids.Themobili・ ヒydifferencebetweenDandD騨alleleswas
slightanditwassometimesdifficulttodis七inguishthe
genotypesDlD,DDorD冒Dl・But二thisproblemcouldbesolvedby
runningthesamplessidebysideonelecrヒrophoresis.The
hamadryasbaboonshadtwoalleles,DandE.Theseallelessee皿ed
tocorrespondtoSandFalleles,respectively,whichBarnicot,
HuehnsandDance(1965)hadfoundinthehamadryasbaboons.The
anubisbaboonshadtheallelesC,D'andE.Thesecorresponded
probablyS,MandFalleles,respec『 ヒively,oftheyellowbaoons
(Papiocynocephalus)asreportedbyBarnicot,HuehnsandDance
(1965),andtotheA,BandCor3,2andユalleles,respectively
describedbyBueヒtner-Janusch(1963,1965),totheA,BandC
alleles,respectively,ofchacmababoons(Papioursinus)as
reportedbyMcDermid,VosandDowinq(1973),andtotheS,皿
andFalleles,respectively,oftheguineababoon(Papio2且 些)
of]」ucotteandGuillon(1979).
Severalresearchersinves七iga'ヒedtheTflocusof
Chaeropirヒhecusbaboons,but,withtheexcep'ヒionofBarnicot,
etal.(1965),theycouldno『 ヒfindtheallelecorrespond-
ingtoourDalleleinthehamadryasbaboons・Also,inrヒhepresent
work,theDallelecouldnoヒbefoundintheanubispopulationsAK
andAI.T'heauthorfoundtheDallelewithalowfrequencyinAW
group,butitwasprobablethatthisgrouphadbeenhybridized
withthehamadryasasdescribedinMaterials.工twasinferred
thatrヒheDalleledidnotexistintheanubisandthattheDl
一8一
alleユedidno・ ヒexistinthehamadryas,becausethefrequency
ofD'allelewasveryhighintheanubisqroupsAK,A工andAW,
butverylowinthehamadτyasgroupsHBA,HBB,HS,HCandHAG。
CrawfordandO'Rourke(1978)reportedthattherewe■eonlytwo
allelesinthehamadryasbaboonshousedattheSukhumiCenter,
USSR.TheD'alユeleinEBA,HBB,1{S,HCandHAGwouldthusbe
theresultofthemigτationofanubisqenesfromoutside・
Pi(Plasmaproteaseinhibito■or(X1-an七itrypsin)=
Theau七horfoundthreeallelesandfourgeno七ypesatthis
locus(Fiq・2)・Astheanubispopulationswerecompletely
monomorphicandthevariantswereobservedintheha皿adryasand
hybridpopulationsonly,itwasinferredthatthevariantallele
Nandghadoriqinatedfromthehamadryaspopula七ions.McDermid,
▽osandDowining(1973)reportedtwoallelesinthechacma
baboons,bu・ ヒtheauthorcouldnotcomparethemobilitiesofrヒheir
alleleswithours.
PA-2(Plasmaprealbumin-2)3
Barnicot,HuehnsandEade(1965)reportedthattherewere
twoallelesatthislocus,butthepresentaurヒhorfoundthree
allelesand6phenotypes.Thephenotypeswiththefast,middle,
slowanddoublebandswereass㎜edtocorrespondtotheqenotypes,
1-1,2-2,3-3,2-1,3-land3-2,respectively(Fiq・2)・The2
allelewasobservedtohavehighfrequenciesinAKandA工popula-
tionswhichwereconsideredtobepureanubis,whiletheother
populationsdisplayedpolymorphism.Thefrequencyofthelallele
wasabout75竃in七hehamadryasgroupsHBA,HBB,HS,HCandHAG・
Thislocusisthusconsideredasaneffectivelocusfor
一9一
discriminatingbe㌻weentheanubisandthehamadryaspopulations.
ES(PlaSmaesteraSe)3
ThisplasmaesterasewasconsideredtoexistintheA20ne
ofBarnicot,HuehnsandEade(1965) .Theaurヒhor-foundtwoes・ ヒ{…rase
bandS:intheplasmaprealbumエnaエeabyusingtheShawandP=asad's
(1970)me七hod.Thephenotypeswiththefast,slowanddouble
bandswereconsideredasexpressionsof土hegeno=ヒypesF♂S-and
FS'respectively《Fig・2)。TheSallelewasapParen七lyatfixa-
tionintheAKandA工populations(consideredtobepureanubis)r
bu'ヒthehamadryaspopulationsexhibi七edhighpolymorphismswith
aboutfifty-fiftygenefrequencies.Thislocuswasalsoan
effec七ivelocusfordiscrimina七ingbe七weentheanubisand
hamadryaspopulations.
6PGD(Redcell6-phosphogluconatedehydrogenase):
Theauthorfoundavariantbandwi七haslight工yslower
mobilitythantheco㎜onbands.Theco㎜onphenotypebeingE,
thevariantphenotypeswerenamedasEFandF(Fig.2).
McDermid,VosandDawninq(1973)alsofoundavariantallelein
thechacmababoons,bu'ヒthisvariantalleleseemedtobe
differentfromtheonefoundbythepresentauthorjudginqfrom
theirmobility.Thepureanubisgroupsdidnothavethevariant,
butthehybridandhamadryasgroupswerepolymorphic.
PGM(Redcellphosphoglucomutase)3
1tmightbethoughttha'ヒthel迦 ⊇PGMwerecontrolledby
twoqeneticloci,PGM一 工andPGM一 工工,asinotherprimatespecies
(Shotake,OhkuraandNozawa,1975)・However,theisozymes
一10一
controlユedbythePGM-IandPGM一 工1ユociapPearedatrathermore
separatepositionsthanthoseofotherprimatesonelectrophoresis.
TheauthorfoundthatthePGM一 工locuswaspolymorphicintheanubis
baboon,while-「 ヒhePG14r工 工locuswa5variabl6even♂in七hehamadryas
baboon(Fig・2)
ADA(Redcelladenosinedeaminase)3
Theco㎜onADAtypeseemedtobecontrolledbythenormal
allele2andthevarianttypewithafasterbandbytheotherへaユ ユele4(Fig・2》 ・TheanubispopulationfromKenya,AK,was
polymorphic.at七his・locus.Theauthorfoundsomevariant
allelesinHAG,HSandHCgroupsanditwaspostulatedtha七the
variantshadbeenbrouqhtintothesegroupsbyi㎜igrants
carryingtheanubisgenes.
CA-1(Redcellcarbonicanhydrase)3
Thevariantallelefoundbythepresen七authorwasconsidered
tobeiden七icalwiththeballelefoundbyBarnicot,JOllyand.Eade
(1967)(Fig.2).・Astheとllele.bwasob6ervedinAK,AWandHAG-
populξLtions,thevarianゼ とouldhaveく)ri官inatedfrom.theanubis
populations.
AK(Redcelladenyla七ekinase)=
Avariantat二thislocuswasobservedinfourhamadryasand
oneanubisgroups.Wepostulatedtharヒthisvariantwasa2-l
heterozyqote(Fig.2).
Wecouldnotfindanyremarkablepolymorphismatthered6ell
enzymeloci.T'heTf,PA-2andPlasmaEsloci,however,were
一1工 一
usefulfortbgeneticalユydifferentiatingbaboonpopula七ions.
Inthatrespect,i七wasfoundthatthegenusPapiodifferedfrom
thegeneraMacaca(Nozawae七aユ.,19753Nozawaetal.,ユ977,
Shotake,1979),Cercopithecus(Kawamoto,unpublish6ddata)and
Theroウithecus(Shotake,1980).,1
Esrヒimarヒionsofhybridizationrate
Gene七icalestimationofthehybridiza'ヒionratebetweenthe
-aロubisand」 蹴yaswas-one-of.themosti批eresUngProblems.
Inconsideringthis,theTf工ocuswasthebes七geneticmarker,
becausetheanubishadtheC,DlandEalleleswhilethehamadryas
hadtheDandEalleles(Table3).The -hybridiza・ ヒionratecanbe
calculatedbythefollowingformula,
qユ ーqo
胚=-rτ 一(G'assandエ'"953)
where皿i・thehybridi・a七i。nrat・'q。th・frequ・n・y。fD'alle1・
inthen。n-hybridi・edp・pulati。nwhichwas・er。'qlthefrequency
ofD蟹alleleinthehybridizedpopulation,andQthefrequency
ofD冨alleleintheanubispopulation.Thehybridizationra七es
oftheMGandMKpopulationswerecalculatedas
O●6667-OMMG=0
.8846.0=0・7537and
O●2907-0
㌔=0.8846-0=0・3286'
respectively3thatis,itcanbeconsideredthat75・4亀and32.9
暑oftheqenesintheNGandMKpopulations,respecrヒively,have
origina七edfromtheanubis・Here,weused・ ヒothevalueofO.8846for
Q.ThatisbecauseBuettner-Janusch(1963,1965)裏eportヒed
theexistenceofanal工elecorrespondingtoourEallelein
KenyaandbecausetheA工populationinourstudycontained7亀
Eand4亀Calleユes.Theestimatedhybridizationratesofthe
一12一
otherpopulationsfromtheanubispopulationaregiveninTable
4.工naddition,thehybridiza七ionratewasestimatedusingthe
polymorphismsatPA-2andEslociandcε 〔lculatingtheaverage
rateofhybridiza七ionfortheTf,PA-2andEsユoci(Table4).
工nthesecases,thevaluesofO.2394andO.6083wereassiqnedto
q。f。rthe2alユeleatthePA-2andtheSalleleattheEsユ 。ci'
respectiveユy,thesebeingtheaverageamongthe5hamadryas
populatio虹s-。 」』5.土he£Ls土i皿ationofahybrid.i.zationrat£-forAW
groupbytheDロallelewasverydifficult,thehybridization
ratewasestimatedbythepenetrationrarヒeofDalleleintoAW
groupinstead・Inthiscase,ΩwasassignedavalueofO.1538,
thisbeingthefrequencyoftheDalleleinHBAgroup(apparently
thepures七hamadryasgroup》.Thepene七ra七ionratefortheAW
groupwascalculatedas
O●0385-OHAW=
0.1538-0=0・2567'
andl-0・2567=0.7433isgi▽eninTable4asthehybridization
rateofAWgroup。
Ωuantificarヒionofeneticvariabilitwi七hinoulation
Thegeneticvariabi工itiesofeachpopulationarealsoshown
inTable4・Theproporヒionofpolyπ10rphicユoci,Pofthepoly'
MG,MK,HS,HCandHAGgroupswhichwereinvadedmoreorless
byanubisgeneswerehiqherthanthoseofthethreeanubis
groupsAK,A■andAW.Theaverageheterozygosiー ヒyperindividual,コH,inthepureanubisqroupsAIくandA工waslessthanO・03・This
issmallerthanthoseofthehamadryasgroups,butlargerthan
those(H繋0・Ol3)ofgeladababoons・(Theropithecusqelada),in
thecentralEthiopianhighland(Sho七ake,ユ980)。 工tisnatural
thatthe.geneticvariabili一 ヒy-of-the-AKgroupsshouldbesInal1,
一13一
sincethisgrouphasbeenkeptinanenclosurefor20yearsago.のTheHvalueofthehamadryasqroupsHBAandHBB(apparen七ly
almostpurehamadryas)wasO.04070naverage,andthisishigher
thanthatfortheanubis(AKandA工).ThefactthattheMGand コ盟1くgroupshadthehighestHvalues,090526andO.0529,respec七ively,
mighrヒsugges'ヒthatthehybridizationwassecondaryintergrada-
tion.Generallyspeaking,thegeneticvariabilityofthegenus
Papioestimatedinthepresentworkwasratherユessthanthose
forthegenus塾4acaca(Nozawae七al.,1977).
Estimationofqeneticdistancebe七weenoulations
Thegeneticsimilarityandgeneticdis七ancebe七weenthe
populationsofbaboonswerecalculatedbytheNei書s(1975)
method.皿atricesofgeneticsimilarityandgeneticdistanceare
presen七edinTable5.Theaveraqegeneticdistancebetweenthe
anubisgroupswasO.0041andthatbetweenthehamadryasgroups
wasO.0024.Thesevaluesapproximatethosebetweentroopξ30f
Japanesemacaque(NozawaetaL,ユ975)andbetweenherdsof
geladababoons(Shotake,1980).Theaveraqegeneticdistance
betweentheanubisandhamadryasgroupswascalculatedas
O.0451.Thisvaluewassliqhtlyhiqherthanthevaluebetween
Japanesemainlandmacaque(Macacafuscatafuscata)andJapanese
Yakumacaque(Macacafuscatayakui),andwasonlyathirdof
theaveragegeneticdistancebetweenspeciesofthegenus
Macaca,whichwasO.1385(Nozawaetal.,1977).However.the
geneticdistancebetweentheanubisandha皿adryasbaboons
estimatedinthisworkisprobablyanunderestimate,because
mos'ヒofthebaboonpopulationsfromwhichtheauthorcol工ected
bloodsampleswereconsideredtocontaingenesfrombo七hthe
anubisandhamadryasbaboons・Theauthor,then,atもemptedtbesti皿a七e
一14一
gene七icdis七ancebe'ヒweenthepureanubisandhamadryasbaboons
byHei,s(1972)formulawhichcorrectsthegene七icdistance
be七weenthehybridpopulationsbythehybridizationrate・That
is,
」x。(・-q)2-2P(ユ ーP)P2Jx
JxY。=(1-P-q)'2-q(1-q)(1-P-q+2pq)-P(1-P》JxY
JY。q2-2(1-P)q(1-P)2」Y
⊥
and■ 。-」kY。/(JX。JY。)2'D。=一 ・n・。'
wherepistheproporヒionofgenesinpopulationXthatcomes
frompopulationY,andqtheproportionofgenesinYthatcomes
f「 。mX'andJXO'JXY。and」YOaretheid・ntity・fgene・c。rres-
P・ndingt。 」X'」XYandJY'・espectiv・ ユy'b・f・rehyb・idi・ati。n
。ccurred・10andDOareth・qen・ticsimilarityanddi・tance'
respectively,beforehybridization.Thecorrectedgenetic
distancesbetweentheanubisandhamadryaspopuユationsaregiven
inparen七hesesinTable5.Theaveragecorrectedgenetic
distancebe・ ヒweentheanubisandhamadryasqroupswascalculated
asbeingO.0679.Thisvaluecoincidedwellwiththa七 〇btainedgbetweenthe四pure闘speciesinferredinourpreviousreport
(Shotake,NozawaandTanabe,1977).
Thegene'ヒicdistancesbetweenthehybridpopulationNGand
theanubisgroupsweresmallerthanthosebetweentheMGand
thehamadryasgroups,whereasthegeneticdistancesbe七weenthe
hybridpopulationMKandthehamadryasgroupsweresmallerthan
thosebetweentheMKandtheanubisgroup.Figure3isthe
dendroqramdrawnfromthegeneticdistancematrixofTable5by
usingtheunweighted-pair-groupmethod(SokalandSneath,1963).
ItcanbeconcludedfromFig・3tha七theMGpopulationhada
cエoserrelationshipwiththearlubisthanwi七hthehamadryas
一15一
baboon,whereastheMIくpopulationhadcloserrelationship壷ith
thehamadryasbaboon.Thisviewissupportedbythecalculated
hybridizationrates(Tabユe4).TheAWpopulationwascloserto
theanubisthanthe瓦Gpopulation,thoughAWpopulationhada
hybridiza『 ヒionrateequaltothatof塾4Gpopulation(Table4).
Correlarヒionsbe・ ヒweenmorholoicalandeneticalhbridindices
andbe七weenrankordersofeneticalindexandbehaviour.
Sugawa■a(1979,inpress)carriedoutanin七ensiveecologi.
calandsociologicalsurveyontheMGandMKgroupsanddeter-
minedamorphologicalhybridindexforeachindividualbelonging
tothesehybridqroups.エattemptedtotestthecorrelation
be'ヒweenthegeneticalindexmeasuredbytheproteinmarkersand
bothofthemorpholoqicalindexandtherankorderofbehaviour
gi▽enbySugawara(1979,inpress)inordertoinvestigatea
detailedstatusofhybridgroup。Table6showsthemorphologi-
calandgeneticalhybridindicesandtherankorderofbehaviour
ineachindividualfromwhichbloodsampleswascollected.The
morphologicalhybridindexwasdeterminedbySuqawara(ユ979)as
follows:eigh『 ヒmorphologicalcharacterswithaclearlydifferen七
appearanceinanubisandhamadryasmaleswereselected。Each
characterwasgivenscoreOforhamadryasappearance,scorel
forintermediateapPearanceandscore2foranubisapPearance・
Themorphologicalhybridindexofeachmaleisthesumofthese
eiqhtscores.TherankorderofmaleswasdefinedbySugawara
(1979)asthearrangementaccordingtQtheirtendencytoherd
females.Thegeneticalhybridindexwascalculatedbythefollow-
inqprocedure・Threeloci(Tf,PA-2andEs)fordis-
criminatingbe七weentheanubisandthehamadryaswereused.Each
alleleattheselociwasbasicallygivenscoreOforhamadryas
一16一
typeandscoヱelforanubistypebutsomecorrectionswere
addedforallelefrequencies,fortheTflocus,allelesD',C,
EandDweregivenscoresl,1,0.landO,respectively,
becausetheanubispopulationshadlOiきEallele3forEslocus,
allelesSandFweregivenscoresO.5andO,respectively,
because50毫Sallelewasexistinginhamadryaspopulations=for
thePA-210cus,alleles2,land3wereqivenO.9,0andO,res-
pectively,becauselO審2aユ ユe■ewasexsit.ing-ilLユ 戯ya」s.-popu■a,
tions.Thegeneticalhybridindexofeachmaleisthesumof
threegenotypescorescalculatedfromtheallelescores(Table6)・
Thecorrelationcoefficientsbetweenthethreeloci,betweenthe
gene『 ヒicalandmorphologicalindicesandbetweentherankorders
ofgeneticalindexandbehaviouraregiveninTlable6.The
correla七ioncoefficien七samongthethreelociwereapproxima'ヒely
zero,sugqes'ヒinqthattherewerenolinkaqedisequilibriumin
thesepopulationsandthatthesepopulationshadpassedthrough
manyqenerationssincethebeqinningofhybridization.The
c6rrela『 ヒiQncoefficien'ヒsbetweengeneticalandmorphological
hybridindicesshowedrelativelyhighvaluesforeachlocus
thoughstaー ヒisrヒicallyno七significanrヒin厘Ggroup,butnon-signifi-
cantneqativevaluesinMKgroup.Also,rankcorrela七ions
berヒweenrankordersofgeneticalindexandbehaviourinMGgroup
showedrelativelyhighervaluethoughs七arヒis七icallynotsignifi-
cantthaninM王(group。SeveraloftheanimalsintheMGgroup
appearedtobeanubisindividualswhich皿ayhavei㎜iqrated
direc七lyin'ヒothisgroupfromnearbyanubistroops・ 工fso,i七
islikely七hattheseanimalshaveelevatedthecorrelations
coefficientsbetweenthegene七icalandmorphologicalhybrid
indicesandbetweenrankordersofqeneticalindexandbehaviour.
一17一
Also,thisphenomenonmaysuggestthattheMGgroupisthe
anubispopulationwhichhasmorerecen七lybeenhybridizedbythe
geneflowfrom'ヒhehamadryasbaboons.
Mira・ ヒionratesamonheihbourinoulations
Next,themigrationratebetweenneighbourinqtroopsofthe
hamadryas,theanubisandthehybridwereestimated.Wright
(1965)derivedaformulaqivingthefunc・ ヒionalrelationships
b・tweenFST(inbreeding・ 。・ffi・ientduet。P。pul・ti。n・ubdivi・i・n)
andm(miqra'ヒionratepergenera・ ヒion)ontheassump七ionof
islandmodelofpopulationstructure,namely,
2.一(1-m)2FST=か
q/q(1噛q)=2N.(2N-、)(、.m)2
2whereqand(ク ㌧standforthemeanandvariance,respectively,
q
offrequenciesofanallelea七avariablelocusamongpopulations,
andNistheeffectivesizeofsubpopulation.Fromthisformula,
wecanestimatetheaveragemiqrarヒionrate(mperqenera'ヒion)by
thefollowinqequation:
2NF
温=1_ST
(2N-1)FST+1
Theauthorcouldestilnateapproximatelytheeffectivepopulation
sizeofhamadryasbaboonsasahalfofitscensussizebythe
followinqprocedure.Nozawa(1972)devisedasimpleformulaby
whichtheeffec七ivepopula『 ヒionsizeofJapanesemonkeytroops
couldbeestima『 ヒed.T'hatis,
2N。(Nc隔1)
N=
琴k孟i・Nl/Nf-Nc■
一18一
whereNi・theeffectiv・p・pul・ 七i。n・i・e'N。 七hecen・u・n㎜ber
。fp・pu…i・n'Nfthecen・u・-ber。f・du・ ・f・m・ ・esandζk孟i
thesumofsquaresofthenumberofgama'ヒescontributedbythe
individualmaleparents。Assumingtha七rヒhisformulaisapPli-
cab・ ・t。 ・h・b・b。 。np。pu…i。n・,wecan…im… ζk蓋iinth・
hamadryaspopulationbyexaminingthecompositionof七roopsor
bands,becauseahamadryastrooporbandconsisrヒsofa
numberofonemaleuni七s(Kummer,1968).Theau土 士Lorcould
examinethecompositionofI{BAandHBBbandsbythefield
・urv・yin・976・ndca・ ・u・・t・dtheapPr・xim・t・ 苧k孟iv・ ・uesby
thenumberofonema工eunitsandthen血beroffemalesbelonq-
inqtothemasshowninTable7,assuminqtharヒaleadermale
hasanequalchancetocontributeoffspringtothenextgenera-
tionbymatingwithhisaccompanyingfemalesinhisownunit.
Asaresult,itwasfoundthattheeffectivepopulationsize
wasapproximarヒelyahalfofthecensussizeinthehamadryas
band.Kummer(1968)descヱibedthattheproportionofadult
malesandfemaleswereonaveragel8.gand32.4老,respectively,
ofthecensusnumberofthehamadryaspopulations.The
effectivepopula七ionsizecanberoughlyestimatedusing
thesevalue・andth・Wrighゼ ・f。rmul・(1938)'N=4NmNf/(Nm+Nf)'
thati・'N=4xO・18N。xO・32N。/(0・18N。+0・32N。)冨0・46Nc-
Thus,wecanaqainsaythattheeffectivepopulationsizeof
thehamadryasbaboonsisapProximatelyahalfofthecensus
size・Also,wecouldes七ima'ヒetheeffec'ヒivepopulationsize
ofanubisbaboonsasonethirdofitscensussizeaccordingto
Nozawa's(1972)me七hod,becausethesocialstructureofanubis
baboonsissimilartothatofJapanesemacaques,Macacafuscata,
一19一
andassumethattheeffectivepopularヒionsizeofhybridqroups
wasavaluebetweenonethirdandahalfoftheircensussize
allocatingpropor『 ヒionallytheirhybridizationrates.
Theaverageeffectivesize(N)oftwoorthreeneighbour-
ingtroopscanbeobtainedfrom・ ヒhehar皿onicmeansoftheN雪s.
Table8qivestheresul七sofestima七ionofmigrationrate
betweenneighbouringbandsandtroopsofhamadryasbaboonsand
ユ⊃etwe.enneighbouringtroopsofa箆ubisandhybridbaboons.Here,
theco㎜onestallelefrequencyateachlocuswasusedforq.
Theaveragemigrationrate(m)betweenAandBbandsintheHB
troopshowsrヒhehighes七valueandthisiscomparablewiththa'ヒ
obtainedamongthreetroopswhichoccurredbyfission(Sugiyama
l960,Kanol964)fromone七roopofJapanesemacaques,Macaca
fuscata,atTakasakiyama(Nozawae七al.,inpress).Nozawaet
al.(inpress)sugges'ヒedtha'ヒthemvaluesobtainedfromseveral
locishouldbeconsideredasunderestimates,becausethevariance
・f・hec・ ㎜ ・n・ ・・ ・・・…fr・quen・i・ ・(6氏2
q)・ ・nt・in・a・amp・ing
variance.Consequently,itshouldprobablybeconcludedtha七
theneiqhbourinqthreetroopsofhamadryasbaboons,HB,HSand
HC,areexchangingindividualsatamigra七ionrateofrather皿ore
thanl2そperqeneration.Assumingtheqene'ヒicequilibriumin
thespeciesborderandhybridzone,themigrarヒionratesbetween
thetwoneighbouringgroupsMGandMKandbe・ ヒweenhybridqroup
NGandanubistroopA工wereroughlyes『 ヒimatedasaboutO.0750
andO.Ol39pergeneration,respecrヒive工y(Table8).
一20一
DISCUSS工ON
Geneヒicvariabilirヒy(H3averagehe・ ヒerozyqosityperindivi-
dua1)ofananubisbaboon七roopwasabou七 〇.02andtha七was
smaller七hanO.040btainedinahamadryasbaboontroop(Table4).
Lucot七e(1979)reportedthatheterozyqosities(H)infourbaboon
ロ コspecエes,旦 塑 鰻,IR.anubis,王 ≧.cynocephalusand2
,.hamadryas
wereO.039,0.Ol8,0.028andO.05,respecti▽ely.Olivier,
Buettner-JanuschandBuettner-Janusch(1974)fσ ㎜dtwopolΣ ㎜oτ 一
phismsonlyattheCAlociofolivebaboon,旦 迦anubis,
livingintheLaipikiadistrictofnorthernKenya.工tisa
somewha『 ヒsurprisingthatdes⇒i・ ヒetheanubisbaboons,王 辿anubis,
havethewidestdistributionofthegenus旦 些,theyhavemain-
tainedarelativelysmallervariabilitycomparedwiththemore
peripheralspecies.Nozawa,Sho七akeandOhkura(1975)andNozawa
etal.(1975)suggesrヒedthatgeneticdriftcouldbeacauseof
lowgeneticvariability(H=0.Ol4)within,andmarkeddifferen-
tiarヒionbetween,troopsoftheJapanesemacaque,1旺acacafuscata・
Sho七ake(1980)alsoconsideredthatbottle-neckeffectwould
diminishthegeneticvariabilityofqeladababoon,Theropithecus .
qelada,(H=0。Ol3)asaresul'ヒofareductionofthepopulation.
Geneticdriftorbottle-neckeffec七maybeinvokedtointerpret
lowgene・ ヒicvariabilityof七heanubisbaboons,sincetheirpopula-
tionstructureissi皿ilaエtothatofJapanesemacaques(Kawaiand
Sugawara,1976a,b)and王 くawaiandSugawara(1976a,b)sugqest
thatthisspecieshasrapidlyexpandeditsdistributionareaand
popularヒionsize.But,thereisatpresentnoevidenceavailable
totestthevalidityofthishypothesis.
Mostofthebaboonpopulationsfromwhichtheauthor
一21一
collectedbloodsa皿pleswereconsideredtocon七ainqenesfrom
bo'ヒhtheanubis奪ndhamadryasbaboons(i.e.,mos七 〇fthemwere
moreorlesshybridpopulations).Thehabi七atoftheHAGtroops
wasmorethan70kmawayfromthehybridzonedescribedbyNaqel
(1973)intheAwashValley-TheHCandHStroopsalsolivedin
thegalleryforestalongtheCassemriverlocatedmorethan30
kmawayfromrヒheAwashValley,whiletheHBAandHBBbandslived
30kmawayfromthewesternedgeofthevalley.Thedis七ribu'ヒion
areaofthesepopulationswerelocatedintheareawhichwas
indicatedasthehabitatforpurehamadヱyasbyNage1(1973).
TheAWtroopwaslocatedl5kmawayfromtheGaribaldipass
whichNagel(1973)indicatedasthewes'ヒernedqeofhybridzone・
Thissuggeststha'ヒthehybridzonebetweenP.anubisandP.
hamadryasisno七asnarrowasoriginallythought・Fieldsurveys
undertakenbyKawaiandSugawara(1976a,b)andbyエwamoto(1980)・
insouth-eas『 ヒEthiopialedthemtoarquethatthehybridzone
ext。nd。d。veranarea。f。b。ut300x3・ ・ 血2,inc・uding七h・
borderbe'ヒweenEthiopiaandSomalia・
工nl976,weobservedthattwohamadryasadult皿aleshad
penetratedirltotheanubistroopAエandmatedwi七hseveralanubis
females.Therefore,weexpectedinl979expeditionthatwe
wouldfindmanyhamadryasgenesthere3however,wecouldfind
fewtypicalhamadryasgenessuchasDandFallelesattheTf
andP-Esloci,respectively,andcouldno'ヒcomple『 ヒelyprovethat
geneflowfromhamadryastoanubishadoccurredinthishybridiza-
tionzoneaswe⊥lasthatfromanubistohamadryas。However,as
wefoundEandlallelesattheTfandPA-210ci,respectively,
whichmiqhtcomefromthehamadryas,thepossibilityofdirect
geneflowfromhamadryas・ ヒoanubiswasnotnecessarilyexcluded.
Althoughgeneflowaroundtheborderbetweenanubisandhamadryas
一22一
habitatswouldmainlybecausedbythemigrationoftheanubis
femaleskidnappedbythehamadryasmalesintothehalnadryastroops
aspostulatedbyNagel(1973),wecouldno七explaintheformation
of『 ヒheMGgroupwhichconsistedofalmos七pureanubisqenesonly
byonewaygeneflow.工tseemsmorelikelythatthehybridization
ofanubistroopsbythehamadryashavebeenaccomplishedgradually
byhybridbaboonswhocouldadapttotheenvironmentofanubis
baboonsInoreeasily.Al七houqh.wecQnsideエthatthespeed.-of⊥ 口・f⊥■-
trationofhamadryasqeneisslowertharltha『 ヒofanubisgenes,our
surveyssuggest七hathybridindividualsofeveエygradebe'ヒweenthe
twospeciesofbaboonsarefoundinourstudyarea・Then,i七
appearsthatthehybridzonebetweentheanubisandhamadryas
baboonsisstiユlexpandingnow.Al『 ヒhoughNagel(1973)arguedtha七
arlevolutionaryapproachtothespeciesproblemshouldconsider
reproductiveisolationbetweenpopulationsintermsofno七 〇nlyinrヒer一
と)reeding(Mayr冒s(1963)defini七ion),bu・ ヒgeneflow(Bigelow曹s(1965)
definition),SzalayandDelson(1979)descエibedbyonlyMayrlsdefi-
nitionwithoutconsideringgeneflowthat璽anubisandl≧ ・
hamadryas ,weresubspecieswithinaspecies.Ourresultsshowthat
thereisnoreproductiveisolationbetweenthetwobaboonsspecies
andthishybridizationphenomenonsuggestsbyBigelow買s(1965)
defini・ ヒionthattheyprobablyaresubspecieswithinaspecies.
Nei(1975)hasshowntharヒthegeneヒicdis七ancebetween
localraceswithinspecieswasintherange、ofO～0.0058,and
betweenspecieswithingeneraintherangeofO.5ハ ノ2.5.Our
estima七ionofgeneticdistancebetweenlocalpopulationsofthe
hamadryasandoftheanubisapproxima七edtotheNei冒srangeof
distancebeヒweenlocalraces.However,theaveraqegenetic
distancebetweentheanubisandhamadryascorrectedfor
一23一
hybridizationratewasO.0679andthisvaluewaslessthanone
tenthdfthedistancebetweenspecies.Thisvaluewasabourヒa
halfoftheaverageqeneticdis七ancebet二weenspeciesofthegenus
Macaca(namely,0.1385:Nozawae・ ヒal.,1977).1」ucotte(1979)
calcula七edtheqeneticsimilari七yamongfour迦 ≧species,
堕 塑 ≧,]≧.anubis,E.cynocephalusandii≧.hamadryasby
usinql410ciandgotvaluesofO.94betweenP・hamadryasand
theotherthreespeciesandaboutO.96amonqthethreespecies
exceptP.hamadryas.Gene七icdistances(0.062andO.040,
respectively)estimatedfromthesevaluesdroppedbelowthe
lowerlilnirヒoftheNeilsgeneticdis'ヒancebetweensubspecies。
Weesrヒimatedtheaveragegene・ ヒicdistanceamongthreeqeneraof
Cercopithecidae,旦 些1,Macacaand⊆iercopithecus,asaboutO・65
(Kawamotoetal.,1977,Shotakel978)andShotake(unpublished
data)es'ヒimateditasaboutO.60betweenTheropithecusandPapio.
Thesevalueswereclosetothelowerlimitofthedistance
be・ ヒweenspecies.Wehadalreadysugqestedthattheclassifica-
tionofAsianmacaquesshouldbereconstructedbylowering
so-calledgenus『 ヒospeciesrankandso-calledspeciesto
subspeciesrank(Nozawaetal・,1977).Here,again,wehaveto
supPortGrove,s(1972),Brettetal.ls(1976)andSzalayand
Delsonls(1979)suqqestionsthat七hesubgenusChaeropithecus
couldbeconsideredasasinqlespeciesbearingthespeciesname
PapiohamadryasLinnaeusl758.On'ヒheotherhand,analternative
opinionistha七thehamadryasisonespecies,Papiohamadryas,
コ ロ(KawaiandSugawara,1976a,b3工wamatQ'1980)ando『 ヒherPap■ospec]・es
.shouldbeluEIpedintoanother層species,Papiocynocephalus.(De▽ore
andWashburn,19633Buettner-Janusch,19663Jol!y,1966ラKingdon,
『1971).Thisopinionisbasedlargelyonthemorpholoqical
一24一
コdiff6r6ncesbetweentheadul七maleSandonrヒhedifferehces■nso-
eialstrudture.between-th。hamadryaa・and土h・ つther・P・Pi。"specie・'璽 ・
Thegene七icdistanceofO.0679_§u99et6.that'.a知ive「 豊ence
timebe・ ヒweenthehamadryasandanubis .can1)ecdlculatedas339'500
years(usingNei騨sμ975]me七hod).Sincethedivergencetime
・・tim・ 七・dfr。mg・neti・di。tance。inth。A。i。nmacaquesw…e1・tive一皇,
1y・ ・mp・ ・ab1・ やP・1ae・n七 ・1・gicaエqsti皿 ・t舌 。n(N・ ・awaeセ ・毛・'1977)'
weinferthatapopulationoftheco㎜onancestorsofthepapio
baboonswerecutoffinadryareainthenorth-eas七ernpartof
AfricaduringthemiddlePleistoceneandwereexposedtosuch
intenseselectiodpressureinahotanddryenvironmenttha七their、morpho工ogyandbehaviourdifferentiatedratherrapidly.
ConsiderinqthatthemorpholρgicaldifferencesbetweenNegroid
andMongoユoidpeoplesinHomosapienshavebeenestablishedonly
forabou七120,000years(Nei,1975),ourconゴectureisnotso
unreasonable.工fthefossilsofPapiosp.founda七Hadar
(JohansonandTaieb,ユ976)andOmo(Eck,ユ977andIwamoto,
personalco㎜unication)inEthiopiaprovetobefromtheco㎜on
ancestorsofthepresent迦 ≧species,itwouldlendconsider-
ablesupporttoourhypothesis.
Sugawara(1979,inpreξ3$)reportedtha七therewere
significantcorrelationsbe層 ヒweentherankorderwithwhichmales
tendedtoherdfemalesandtheirmorphologicalrankorder
.(hybridindices)fromhamadryastoanubisinbo七hthehybrid
qroupsMKandMGandsugges・ ヒedthatthebehaviourofhamadryas
adultmaleswascontrolledbyageneticfactor.工ncontrast,
therearefewcorrelationsbe七weenmo∫phologicalandgenetical
indicesinthepresentanaユyses,perhapsbecausethegenetical
indexiscalculat二edonlyfromthreemajorgenesasmarkersand
一25山
thesemaゴorqenesmayhavecontinuedtoseqregateandrecombine
independentlyofthemorphologicalcharactersthrouqhseveralqen-
erationsafterhybridizarヒion.Moreover,wecannotidenrヒifyany
significantcorrelationsbetweenthebehaviourofhamadryasadult
malesandtheirpossessionofthethreemajorgenes.工tseems
likelythatthe」Lackofthesecorrela・ ヒionsarebecausethemorpho-
loqicalandbehaviouralcharac七ersobservedbySugawaraarecon-
trolledbypoユygenicfactoユ ニs-segエ.egatinqindependentlywitheach
other.
Olivier,Buettner-JanuschandBuettner-Janusch(1974)
attempー ヒedtoestimatethemigrationrateperqeneration(m)
amonqninetroopsofanubisbaboonsinhabitingin工Iaipikia
districtinnorthernKenyabyusinqtwopolymorphiclociof
carbonicanhydraseasmarkers.Theyes七ima'ヒedmattherange
fromO.094toO.214correspondingtoN(effectivepopula七ion
size)varyinqfrom85to30,respectively.Theauthor's
estimateofaveraqemiqrationra七e(m)amongthreetroopsof
hamadryasbaboonswasO.12andthiswouldseemtobecomparable
tothoseobtainedamongtheanubistroopsstudiedbyOlivier,
Buettner-JanuschandBuetrヒner-Janusch(1974),thedis七ribution
areaofthetroopsbeingapproxima¶ ヒelyequalandtheeffective
popularヒiollsizesbeinqinasameranqe.Assumingageneヒic
e喧uilibriuminthehybridzone,themigrationra七ebe'ヒweenthe
twoneighbouringhybridgroupMGandMKwasrouqhlyestimarヒedas
aboutO・075pergenerationandthisvaluewasconsideredtobe
compatiblewiththeresul七soffieldobservationsugqestingthat
MGandD4Kgroupshadexchangedatleastafewindividualsduring
thelastthreeyears(Sugawara,inpress).Thus,wecanconclude
tha『 ヒ 七hereismuchmiqrationberヒweentroopsofbaboonsandthat
一26一
themiqrantqenesfromonekindofbaboonhavebeendiffusing
intothepopulationsof・ ヒheotherkinds.
ACKNOWLEDGEMENTS
Theau『 ヒhorwishestoexpresshis・ ヒhankstoDr.KenNozawa
andDr-MasaoKawaiforencouraqingandsupervising『 ヒhiswork。
ThanksareduetoDr.MitsuoIwamoto,Dr.UmeyoMori,Dr一
王{ideyukiOhsawa,Dr。KazuyoshiSugawara,Dr・YuichiOnoand-his
teamatFacultyofScience,1くyushuUniversity,fortheirvaluable
helpduringresearchexpeditionandMr.YoshiIくawamotoandMiss
YoshikoOhkuraandMissAtsukoOhnofortheirtechnical
assistanceandinpreparingthis皿anuscript・Theauthorthanks
alsoDr.Robin工.M.Dunbqrforhisvaluableco㎜en七stothe
Englishexpressionofthismanuscriprヒ.Theauthorthanksalso
Drs.DouqlasObeckandOwenWoodoftheNAMRU5工nstituteofthe
UnitedStatesofAmericaandMr.GuilioTar七agliawhokindly
offeredlabora七 〇ryandfreezerfacilitiesforstoringblood
samples.Theauthorisaユsoindeb七edtothefollowingfortheir
coopera七ion=MessersTechiomeAshine,工ealemBerhanu,Hagos
Yohannes,TaddesseGabre-Michaiel,Ande-BerhanuIくidane,Tesfaye
Wundesa,Akinori⊃4izunoandMikioKajioftheEt二hiopianWild
lifeConserva七ionOrganiza七ionFDr.ShiburTedlaandMrs.Misrak
EliasofAddisAbabaUniversity=HisExcellencyAmbassador
NitsuoHashiz㎜eandHessersToshiakiTakahashi,YujiHaradaand
KuniyoshiOhtaoftheEmbassyofJapantoEthiopia3Messers
SadanoriTaquchiandYasushi工nabaofJapanOverseasCooperation
Volunteer,andthemembersoftheJapanMonkeyCentre・
一27一
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一35一
TablelBloodsamplesofbaboonsinthepresentwork
欲
肛
甜
鵬
猷
冊
冊
田
鵬
肌
(28)
(27)
(13)
《18)
(43)
(39)
(19)
(35)
(49)
(292)
PapioanubisfromKenya
papioanubisof工totroop
PapioanubisofWelenchititroop
HybridofGorgetroop-
HybridofKerrayutroop
PapiohamadryasofBassakaAband
PapiohamadryasofBassakaBband
PapiohamadryasofSabureGorgetroop
PapiohamadryasofCasselnGorgetroop
PapiohamadryasofAwashGameReservetroops
Abbreviation
-凸
2
一
一
b
A
A
l
f
P
P
A
T
y
t
即
肚
晩
恥
㎞
c。
SE昂
SEamSal
P
P
α
β
一
一
㎝
p
乱
皿
肋
恥
I
I
I
D
一
脚
工
G
M
M
A
H
P
G
G
D
P
6
a・工
P
P
A
D
C 1一A
I
A
B
工
D
一
一
一
P
P
H
H
H
A
C
6
D
D
D
C
A
G
M
L
L
SEl
H
D
1
0
D
K
S
e
T
I
A
E
C
Table2 Listofbloodproteinsexamined
Nameofbloodprotein Reference
PlasmaPrealbumin-1
Plasmaprealbumin-2
Plasmaalbumin
Plasmatransferrin
Plasmahaptoglobin
Plasmaproteaseinhibitor
Plasmaslowα2globulin
Plasmaceruloplasmin
Plasmaamylase
Plasmacatalase
Plasmacholineesterase
Plasmaesterase
Plasmaalkaユinephosphatase
Plasmaleucinaminopeptidase
Hemoglobin一 α
Hemoglobin一 β
Cellphosphohexoseisomerase
Ce116-phosphogluconatedehydrogenase
Cellphosphogユucomu七ase-I
Ceユlphosphoglucomutase-II
Celladenosinedeaminase
CellNADH-diaphorase
Cellcarbonicanhydrase-I
Cellcarbonicanhydrase-II
Ceユ ユacidphosphatase
Cellglucose-6-phosphatedehydrogenase
Cellmalatedehydrogena8e
Cell
Cell
Cell
Cell
Ceユl
Cell
Ceユ1
1actatedehydrogenase-A
lactatedehydrogenase-B
tetrazoliumoxidase
isocitratedehydrogenase
adenylatekinase
esteraseD
esterase
Barnicot,HuehnsandDance
Barnico七,HuehnsandDance
Shotake(1979)
Barnicot,HuehnsandDance
Ishimoto(1972}
Omotoetal.に970)
Barnicot,HuehnsandDance
Imlah(1964》
Oqitaeta1.(ユ966)
ShawandPrasad(1970)
Barnicot,HuehnsandDance
Barnicot,HuehnsandDance
ShawandPrasad(1970)
Scandalios(1964》
Ishimo七 ◎,Kuwataand
Ishimoto,Kuwataand
Tashianeta1.
工shimoto
ShawandPrasad
ShotakeandNozawa
Shotake
Shotake
BaurandSchorr
Ishimoto,
Shawand
Hopkinsonetal.(ユ973)
Kitaゴimaetal.(1975)
(1965)
(1965)
(1965)
(1965)
(1965)
(1965)
Shotake(1975)
Shotake(1975》
Shotake,OhkuraandIshimoto(1977)
Ishimoto(ユ972)
Shotake,OhkuraandNozawa(1975》
Shotake,OhkuraandNozawa(、1975)
Spencer,HopkinsonandHarris(1968)
Ishimoto(1972)
Shotake(=L975)
(1972)
(1972)
(1970)
に974)
(1974)
(1974)
(1969)
1くuwataandShQtake(1974)
Prasad(1970)
Table3.
Allelefrequenciesofvariableproteinlociint血
ebaboonpopulations
Locusand
aユlele
τf
コ
D D E C
piP Q N
PA-2
- 凸 2 「 」
PユasmaF
EsS
6PGD
E F
PGM-14
5
PGM-II3
4
ADA
2 4
CA-1
a b
AK- う ●
Baboonpopulation
AKAI
A珂
0 1.0000
0 0
0 0.8846
0.0769
0.0385
0.0385
0.8846
0.0769
0
1●00001●
00001●
0000
000
000
00・01920・
1923
1●00000・
98080・
8077
000
000
1・00001●
00001●
0000
1。00001●
00001●
0000
000
1・00000●
8462
0.1538
0.8462
0.1538
1・00001・
00000.8077
000●
1923
0・67860●
84621●
0000
0●32140●
15380
0●92861●
00000・
9231
0・071400●0769
1・00000●
88461●
0000
00●11540
MGMK
0.1389
0.6667
0.1667
0.0278
0.1628
0.2907
0.5462
0
0.94440・
9884
0.05560
00.0116
0●05560●
6395
0・94440・
3256
00.0349
0●33330.3023
0●66670・
6977
0●97221}。8953
0・02780●
ユ047
0●75000●
9651
0●25000・
0349
0・88891●
0000
0.11110
1●00000・
9884
00.0116
1●00001●
0000
00 1.00001●
0000
00
HBA
HBB
HSHC
HAG
0.1538
0.0257
0.8205
0
0 ♂ Q ♂ - 凸
Ω U 8 2
7 ・ 7 4
0 ∩ U Ω U
● ● ●
ハ U O ハ U O
0.0571
0.0714
0.8714
0
0.1122
0.1122
0.7755
0
0.1182
0.0839
0.7979
0
1●00000.89470.97140●
9898
00●105300
000。
02860●
0102
0.9552
0.0431
0.0017
0.82050●
73530●
74290.8470
0●08970●
17650●
25710●
1224
0●08970●
088200●0306
0.6573
0.2780
0.0647
0●26790・
19440●
54290・
4796
0●73210●
80560・
45710●
5204
0.4739
0。5261
0・88460●
78950・
95710・
94900・
9633
0●11540。
21050・
04290●
05100・
0367
1●00001●
00001.00001.00001●
0000
00000
1・0000ユ
。00001・
00001・
00000●
9913
00000●
0087
1●00001●
00000●
97140●
94900●
9843
000●
02860●
05100●
0157
1●00001。
0000'1・0000ユ
●00000・
9739
00000●
0261
0098721・
00000●
94290●
95920●
9912
0・012800・05710●
04080・
0088
Tabユe4 Quantificationofgeneticvariabilityand
hybridizationofbaboonpopuユations
Ppoly 員=1一 Σq呈ne=1/Σ 強呈
hybridization1)
rate
AK
AI
AW
MG
猷
2/3430●0588
5/3430●1471
5/34塁0●1471
7/34=0●2059
7/34=30●2059
HBA5/34=0●1471
朋B
HS
HC
HAG
5/34
7/34
7/34
7/34
0.1471
0.2059
0.2059
0.2059
0.0167
0.0286
0.0363
0.0526
0.0529
0.0363
0.0451
0.0415
0.0421
0.0471
1.0170
1.0294
1.0377
ユ.0555
1.0559
1.0377
ユ.0472
1.0433
1.0440
1.0494
0.7433
0●7537(0●6ユ79)2)
0●3286《0。2245)
0.0290
0.0314
0.0807
0.1268
0.0948
1)penetrationrateofanubisgenes
2》thefiguresinparenthesesaretheaverageofthreeユoci,
τf,PA-2andEs'
Table5Matrices。fgeneticdistance《D)underU・ediag。nalandgeneticsimilarityに)
ab。veU'・diag。naユb・tweenev・ 可P・ir。fbab。 叩P。pulati。n・.Thefigure。in
parenthesesaregeneticdistancescorrectedbyhybridizationrate.
AK AI AW
AK
A工
AW
0.0024
0●99760・9935
0●00650●0036
0.9964
MG
MK
0●01ユ60・00640・0058
0●03230●029ユ0●0208
翠BA
HBB
HS
HC
HAG
0●05500●04730。0393
《0●0583)《0●0503》 《0・0774)
0・04960・04230・0351
(0●0541)(0●0451)(0●0688)
0●05460●047ユ0。0411
(0●0790》 《0・0546)(0●0933)
0●05460・04790●0406
⊂0●0719)(0・0684)(0.1064)
0●04800●04130●0329
(0・0582》 《0●0505)《0・0830》
MG MK
0●98850●9682
0●99360●9713
0●99420●9794
0.0179
0.9823
0●03670●0034
0・03340●0033
0・03220・0064
0●03080・0038
0・02810・0028
HBA HBB HS HC 胤瓦C
0●9465-0●9516.0●94680●94680●9531
0●95380●95860・95400・95320●9596
0●96ユ50●96550・95970●96020.9676
0●96400●96720●96830●96970●9723
0●99670・99670●99360.99620●9972
0.0011
0●99890●99660・99770・9974
0●00340●0052
0●99480・99590・9962
0・00230.00410.0008
0●99920・9994
0●00260。00380●00060●009
0.9991
Table6Morpholoqica1(M.1.A.)andGeneticalIndices{G・1。A・)of
anubis,BehaviouralRankOrder(B.R.O.)andcorrelation
coefficientsamongthreeqeno七ypesbetweenM.1.A.and
G.1.A.andbetweenB.R.O.andrankorderofM.1.A.
G.工.A.
Name★ M.1.A穿 B.R.O書Tf Es PA-2 Total
Gorgeqroup
Matiou
I、ouis
Maler
Mirabeau
Baboof
Necker
Green
HagOS
Adult6
Adultδ
Adult6
Adultδ
Adultδ
Adult6
Adultδ
Adultδ
6
6
5
4
3
3
2
3
1
1
1
1
1
1
1
6
7
2
8
3
4
5
1
CD.〔2.0}
D。E(1.0)
DD(0)
D.D8(2.0》
D曾D「(2.0)
D曹E(1.1)
DIDI(2.0)
DD(0)
FS(0.5)
SS(1.0)
SS(1.0)
FF(0}
FS(0.5)
FF(0)
SS(1.0)
FF(0)
2-2(1.8》
2-2(1●8)
2-2(1.8)
2-2(1.8)
2-2(1●8)
2-2(1●8)
2-2(1.8》
2-2(1・8)
4.3
3.8
2.8
3.8
4.3
2.9
4.8
1.8
Kerrayugroup
Doro
Feres
Zefen
Igユr
D]oro
Jirat
Kusil
Posta
Yabelo
Stav
Geta
Adultδ
Adultδ
Adultδ
Adultδ
Adultδ
Adultδ
Adultδ
Adultδ
Adultδ
Adultδ
Adultδ
9
8
6
6
6
5
4
3
2
2
2
4
9
3
0
0
6
7
8
2
5
1
1{
1
EDl(ユ.1)
ED(0.1)
ED・(1.1)
DIDI(2.0)
EE(0.2)
D.D冒(2.0)
DD(0)
ED冒(1.1)
ESl(1.1)
DD。(1.0)
EE(0.2)
FS(0.5)
SS(1.0)
FF(0)
FF(0)
FF(0)
FS(0.5)
FS(0.5)
FS(0.5)
SS(1.0)
FS(0.5)
SS(1.0)
1-2(0・9)
1。1(0)
1一 ユ(0)
1-2(0●9)
1-1(0)
2-2(1●8)
2噂2(ユ ●8)
2-2(1.8)
1-2(0・9)
1-1(0)
2卿2(1.8)
2.5
1.1
1.1
2.9
0.2
4.3
2.3
3。4
3.0
ユ.5
3.0
1.Correlationcoefficients
Gorqegroup(n=18)
Tf&Esr=0.0361
Tf&PAr=-0.0616
Es&PAr=0.1344
Kerrayugroup(n冨43)
Tf&Esr=-0.176
Tf&PAr=0.OlO
ES&PAr=0.002
amongthreeqenotypes
P
P
P
<
く
く
-孕-●-ゆ
●
●
●
0
0
0
P
P
P
<
<
<
-白-噸1
・
●
●
0
0
0
II.CorrelationcoefficientsbetweenG.1.A.
Gorgegroup(n=8)
●
●
●
●
-凸2
{5
4・
andM.1.A.inadultmales
r=0・4429
r=0.
r=0・4141
r30●6193
r=-0・3601
r=-0.3663
r=-0。4229
r=-0・3228
P
P
P
P
<
<
<
<
-凸-凸1
10
●
●
●
●
0
0
0
0
P
P
P
P
<
く
く
く
l
1
1卿1
●
・
●
●
0
0
ハ∪
0
III.RankcorrelationsbetweenB.R.0.andrankQrderofG.工.A.
Gorgegroup(n=8)
r80.47620.1〈p
Kerrayugroup(n=1ユ)
rニ ー0.19540.1くp
禽CitedfromSugawara(1979rinpress)。 Fordetailedexplanationseethetext.
Table 7Age-5excomposi七ionsofBassakahamadryastroopandestimationofeffectivepopuユation
昌izeinJanuaryl976・
BandortrOOP Adultm。lesAduユtandSub・dultYounqst。NC
subadultfemalesmales3yearsoユdみ 孟)N(N)・)・N(w)・)■
HB-Aband
HB-Bband
HB-trOOP
14(8)1)
9(5)
23(13)
0
2
2
2
1
3
nO
4
21
8
0
只り
2
2
4
70.
45
1ユ5
4
2
6
ダり
3
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0
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4
2
6
へ」
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4
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5
1)
2}
3}
り
Thefiguresinpare真thesesqivethenumberofleadermaleswithunits
S㎜ofsquareofthenumberoffemalesinonemaleunits
EffectivepopuユationsizeestimatedbyNozawalsformula
EffectivepopuユatiopsizeestimatedbyWrighゼsformula
Table8 Estimati。n。fmig=ati◎n・ate{孟pergene・ati・n)b・tweenneighb◎ringbandsandtr。 。P・
oftheha皿adrya8baboon8andbetweenneighbouringhybτidandanubi6group5
Groupoftroops
(Censussize)
HBAσ0》
HBB{45)
HB(115)
11S(150)
llS(150}
llC{150}
HB(115)
HS(150)
HC(150)
MG
MK
(63)
(85》
AI(60)
MG{63}
Va=iable
lOCUS
Tf
pi
PA-2
P-Es
6PGD
照
Tf
pi
PA。2
P-ES
6PGD
ADA
AK
Tf
pi
Pバ ー2
P-Es
6PGD
ADA
AK
Tf
pi
PA-2P-Es
6PGDADA
AK
Tf
pi
PA-2P-ES
6PGD
PGM-I
PGM-II
ADA
TF
pi
PA-2ロ
P-E8
6PGD
PGM-IPGM-II
ADA
AK
C◎ ㎜onest
allele
E
Pl
S
E
-
E
Pl}S
E
2
1
E
P
ユ
S
E
2
1
E
P
l
S
E
2
1一
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P
2
S
ε
4
3
2
コD
P
2
S
E
4
3
2
1
Allele frequency
q
HBA
0.8205
1.0
0.8205
0.7321
0.8846
0.9872
HB
0.8276
0.9655
0.7946
0.7895
0.8534
1.0
0.9914
HS
O.8714
0.97140.7429
0.5429
0.9571
0.9714
0.9429
HBB
0.8421
0.89470.7353
0.8056
0.7895
1.0
HS
0.8714
0.9714
0.7429
0.45710.9571
0.9714
0.9429
HC
0.7755
0.9898
0.8470
0.4796
0.9490
0.9490
0。9592
HB
0.8276
0.9655
0.7946
0.7895
0.8534
1.00.9914
HS
O.8714
0.9714
0.7429
0.4571
0.9571
0.97140.9429
HC
0.7755
0.9898
0.8470
0.5204
0.9490
0.94900.9592
MG
0.6667
0.9444
0.9444
0.6667
0.9722
0.7500
0.8889
1.0
AI
0.8846
1.0
0.9808
1.0
1.0
0.8462
1.0
0.8462
0。8846
MK
0.2907
0.9884
0.3256
0.6977
0.8953
0.9651
1.0
0.9884
MG
0.6667
0.9444
0.9444
0.6667
0。9722
0.7500
0.8889
1.O
l.0
一q
0.8313
0.9474
0.7779
0.7689
0.83710.9936
0.8495
0.9685
0.7688
0.62330.9053
0.98570.9672
0.82350.9806
0.7950
0.5113
0.9531
0.96020.9511
0.8248
0。9756
0.7948
0.5890
0.9198
0.9735
0.9645
0.4787
0.9664
0.6350
0.6822
0.9338
0.8576
0.9445
0.9942
0.7757
0.9722
0.96260.8334
0.98610.7981
0.9445
0.9231
0.9423
2
qσ
0。000090
0.002778
0.001822
0.001370
0。002247
0。000059
O.002700
0.000005
0・OOO650
0.055200
0.002650
0。000195
0.000600
0.002300
0・000084
0・002705
0。010030
0.000015
0・000126
0.000064
0.001533
0.000100
0,001800
0・020733
0・002200
0●000433
0.000433
0。035350
0・000480
0.095730
0・000240
0。014800
0●011570
0。003090
0.000030
0●01ユ867
0●000777
0.000351
0.027975
00000207
0●002336
0・003064
0.005936
0●003321
Fs「r
0・000642
0.055750
0・0二LO546
0●007710
0・016478
0・004277
0・02ユ127
0。000ユ63
0・003656
09234893
0・030886
0.013830
0●0ユ8868
0.015821
0。004416
0.Ol6597
0。004014
0.000336
0。003293
0●001375
O.OlO609
0。004202
0・011036
0.085682
0・・029810
0.016795
0.012670
Og141L650
0。0ユ4780
0.413030
0・001ユ07
0・023940
0・094740
0.058950
0・000570
0。068201
0・028778
0.009750
0●202132
0●015109
0・014500
0.058473
0・083606
0・061048
FST
0.015901
0.046199
0・006550
0曾024401
0。093596
0・060177
一N
27
65
75
68
一m
0.317・
0.0711
0.293
0.120三
29
21
0.075.
0.138
Lggend
Fig.ユ
Fig.2
Fig.3
Locationofpopulationsf=omwhichthebloodsamples
wereco1ユectedincentraユEthiopia.
Zymogramofeユectrophoreticvariantphenotypesfound
intま 」iswork.
DendrogramdramfromtheNei・sgeneticdistance(D)
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